A New Shark and Ray Fauna from the Middle Miocene of Mazan, Vaucluse

Swiss J Palaeontol (2011) 130:241–258 DOI 10.1007/s13358-011-0025-4

A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) and its importance in interpreting the paleoenvironment of marine deposits in the southern Rhodanian Basin

Nicolas Vialle • Sylvain Adnet • Henri Cappetta

Received: 10 March 2011 / Accepted: 29 August 2011 / Published online: 16 September 2011 Ó Akademie der Naturwissenschaften Schweiz (SCNAT) 2011

Abstract Recent paleontological excavations of the the last decades from several localities in the Natural Park Middle Miocene sandstones (top of the ‘‘Schlier’’ facies) of Lube´ron (South of Vaucluse). Selachian associations near Mazan, Vaucluse (southern France) uncovered a rich recovered in the Middle Miocene of Vaucluse are quite selachian teeth assemblage including 34 shark and ray particular because they indicate either neritic or deepwater species. This great diversity of selachian taxa provides new deposits. Such difference in faunal composition between information about the palaeoenvironmental settings of sites can be interpreted as a difference in age, a geo- some Middle Miocene deposits in the southern Rhodanian graphical heterogeneity of environmental deposits or a basin. For instance, the co-occurrence of some deepwater mixture of both. A new fossil locality, found in 2007 near Squaliformes and Rajidae with numerous Carcharhinifor- Mazan, was collected by one of the authors (H. Cappetta) mes and Myliobatiformes, usually inhabiting the conti- with the kind help of Eric Collier. A preliminary analysis nental shelf, suggests a deepwater deposit inshore, as indicates that the new rich selachian fauna comprises both adjacent to a submarine canyon area. types of the previous associations. The aim of this work is to improve our knowledge about the selachian diversity in Keywords Selachians Middle Miocene Southern the southern Rhodanian basin during the Middle Miocene, France Southern Rhodanian basin Paleoenvironment by listing in detail the selachians from Mazan and com- paring them with material collected in Vaucluse and in neighbouring areas of He´rault. Introduction

The earliest studies on the Miocene fossil selachians from Geological settings Vaucluse were by Fischer (1878); Leriche (1906) and Joleaud (1907, 1912) who published a detailed review of The fossil locality of ‘‘Mazan’’ (Fig. 1) is located at the selachian remains from the southern Rhodanian basin. eight kilometres east of Carpentras (GPS coordinates: These faunas were partially updated from new material N044°03002.800 E005°08015.800) and is mapped as ‘‘Helve- (Cappetta et al. 1967; Ledoux 1972) or from comparative tian’’ (BRGM 1975) formally equivalent to the Langhian– analysis with nearby contemporaneous localities (Cappetta Serravallian period. This age was formerly reported as 1970, 1973). More recently, Brisswalter (2009) published a Vindobonian by Demarcq (1970), a local stage which note on new selachian teeth assemblages collected during corresponds precisely to the lower part of the so-called ‘‘Helvetian’’, and thus, to the Langhian only. The sediment consists of grey sandstone, composed of sorted sand, that N. Vialle S. Adnet (&) H. Cappetta outcrops as a monotonous layer around Carpentras (Joleaud Laboratoire de Pale´ontologie, UMR 5554, Institut des Sciences 1907, 1912). The thickness of this layer varies between 350 de l’Evolution, Universite´ de Montpellier II, Sciences et and 500 m (Demarcq 1970) according to its erosion and Techniques du Languedoc, Cc 064, Place Euge`ne Bataillon, 34095 Montpellier Cedex 5, France location. This unnamed formation probably corresponds to e-mail: [email protected] the upper part of the blue ‘‘Schlier’’marls Formation that 242 N. Vialle et al.

Fig. 1 Left. Map of the major Miocene deposits of Vaucluse with Lithostratigraphic framework of the Rhone–Provence Molassic series, selachian teeth: 1,‘‘Bonpas’’; 2, ‘‘Caumont’’; 3, ‘‘Mazan’’; 4, ‘‘Saint- simplified from Besson et al., 2005. Black: continental surfaces, grey: Didier’’; 5, ‘‘Cabrie`res d’Aigues’’ (redraw after Demarcq, 1970 Bioclastic limestone-dominated deposits (‘‘molasse’’ s.s.), white: modified and Besson et al. 2005 for the Latest Burdigalian network of Sand and marl-dominated deposits. Fl.In: Major Fluvial incisions fluvial valleys: L. Burd fluvial valleys). Right. Early-Middle Miocene occurs in the entire ‘‘Miocene Molasse basin in the Rhone– dried, then treated with a 10% solution of acetic acid over Provence’’ area (Gignoux 1960; Demarcq 1970). Besson a period of 1 or 2 days according to the size and quantity et al. (2005) recently confirmed the Langhian age of these of the residues. After rinsing in clean water and drying, peculiar sandstones and sandy marls, which are sand- the final residues were sorted under a binocular micro- wiched between two major molasse deposits (see Fig. 1). scope. The fossil material consisted exclusively of iso- In these sandstones layers, Joleaud (1907, 1912) and lated selachian teeth, many teeth and vertebral centra of later Cappetta et al. (1967) reported a relatively deepwater Actinopterigians and fragmentary bones of marine ver- shark fauna near ‘‘Bonpas’’, at the SW of our study area. tebrates. The selachian teeth, particularly those of sharks, Closer, the fossil locality of ‘‘Saint-Didier’’ (Ledoux 1972) were often in poor state of preservation with only the exhibits similar sandstone layers and a similar selachian crowns remaining. The smaller teeth, ranging in size fauna with numerous deepwater taxa. These deepwater between 2 and 4 mm were the most complete. The poor faunas are clearly different from those previously recov- preservation of the larger fractions is probably due to ered elsewhere in the Middle Miocene of southern Rho- taphonomic bias. This assumption is supported by danian basin by senior authors (Fischer 1878; Leriche observation that many small teeth are rounded, suggesting 1906; (Joleaud 1907, 1912) but they are consistent with that the material may have been carried on the sea floor recent studies (Besson 2005; Besson et al. 2005) on the before final deposition. It is thus possible that part of dynamics of the ‘‘Rhodano–Provençal’’ basin that observed material collected at ‘‘Mazan’’ was not strictly autoch- that the Middle Miocene has been marked by two maximal tonous and has been transported from nearby. All the transgressions, one dated as Langhian and the second as figured material is housed in the University of Montpellier Serravallian. (Collection number with abbreviation MAZ). The sys- tematics and dental terminology of taxa follow Cappetta (1987, 2006). The studied fossil material represents 25 Systematic palaeontology species of sharks belonging to 22 genera and 9 species of rays shared in 8 genera. Only three genera are recognized A 60 kg sediment sample was sieved through a series of as exclusively fossils: Paraheptranchias, ‘‘Oligodalatias’’ screens, the finest of which was 0.3 mm. Residues were and Carcharoides. A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) 243

Order Hexanchiformes Buen, 1926 well marked like the distolingual depression. A well- Family Heptranchidae Barnard, 1925 developed uvula with small mesial foramina is present on Genus Paraheptranchias Pfeil, 1981 the lingual face (Fig. 2-1b). The infundibulum opens Paraheptranchias repens (Probst, 1879) beneath the uvula, at the level of the well-developed Notidanus repens (Probst, 1879) transverse lingual bulge, and is followed by a more or less Notidanus avenionensis (Joleaud, 1912) marked groove toward the basal edge of the root. The Material: 5 incomplete teeth. upper tooth has a narrower and more erect cusp than the lower teeth. The distal heel is well separated from the main This unusual species is represented by incomplete frag- cusp. The apron is broader at its base. On each side, there ments, primarily of the crown, probably those of lower are two small foramina. On the lingual face there is a small teeth. The root is missing. The main cusp is highly uvula, which is not well marked. Beneath the uvula, there developed with a sigmoid shape. The mesial cutting edge is are two axial foramina followed by a slight groove. limited to the upper part of the cusp. The cusp is flanked by Remarks. The presence of a slightly serrated mesial small denticles, which generally separate the acrone from cutting edge and absence of fold on the uvula are indicative the accessory cusplets. of the living species Centrophorus granulosus. This species Remarks. This peculiar species was described by Probst is widespread in the Miocene sediments of Vaucluse, from (1879) in the early Miocene of Germany and well illus- the Early to Late Miocene (Cappetta et al. 1967; Ledoux trated by Barthelt et al. (1991). Joleaud (1912) also 1972; Cappetta 1975; Brisswalter 2009). reported a unique specimen of P. repens, formerly as Notidanus avenionensis, from the Middle Miocene deposits Genus Deania Jordan & Snyder, 1902 in the region of Avignon, southern France (Cappetta 1970, Deania aff. calceus (Lowe, 1839) 1987). Material: 10 teeth. (Fig. 2-2, 2-3) Family Hexanchidae Gray, 1851 Compared to Centrophorus, lower teeth of Deania are Genus Hexanchus Rafinesque, 1810 more compressed labiolingually and the crown is narrower. Hexanchus sp. There is no serration on the mesial cutting edge. On the Material: 2 teeth. labial face (Fig 2-2a), the apron is very wide and thick, but its limits are slightly marked from the root. On each side of The two teeth are both crown fragments, probably from the apron, there are two well-developed marginal foramina. lower teeth. The crown is easily separated from the pre- The labial depression is well marked. On the lingual face vious taxon by the large number of accessory cones, which (Fig. 2-2b), the uvula is well marked but small and is decrease regularly distally. On one tooth, one can observe a flanked by marginal foramina, which reappear beneath the small serration on the lower half of the mesial cutting edge straight bulge and may be followed by a slightly marked of acrocone. groove. The upper teeth (Fig. 2-3) have a straight and high Remarks. The living species Hexanchus griseus Bon- cusp. The apron is broad with a well-developed marginal naterre, 1788 is known from southern France from the foramen on each side. The crown has both distal and mesial Pliocene in Le-Puget-sur-Argens (Var) (Cappetta and Nolf, heels. Teeth differ from those of Centrophorus by the 1991). It was recently reported in the Langhian–Serraval- reduction of the uvula on the lingual face. The lingual lian of Cabrie`res d’Aigues (Vaucluse) (Brisswalter 2009). transversal bulge is well developed and bears a foramen. At The poor state of preservation is insufficient to confirm its end, there is another small foramen followed by a whether this material belongs to the recent species or not. slightly marked groove. The root is large at the base of the Order Squaliformes Goodrich, 1909 crown and thin at its basal edge. Family Centrophoridae Bleeker, 1859 Remarks. The number and position of the foramina on Genus Centrophorus Mu¨ller & Henle, 1837 the lingual face of these teeth are distinctive of genus Centrophorus aff. granulosus (Bloch & Scheinder, 1801) Centrophorus and indicative of the recent species Deania Material: 42 teeth. (Fig. 2-1) calceus. The remains of this genus are very rare in the fossil record but they have been already observed in the The lower teeth are broad and strongly compressed labio- bathyal deposits of the Lower and Middle Miocene of lingually with a very large cusp pointing to the rear. The Vaucluse (Cappetta et al. 1967; Ledoux 1972). distal heel has a convex shape. On the labial face (Fig. 2- 1a), the apron is quite developed with a base broader than Family Dalatiidae Gray, 1851 its extremity and many small foramina are present on both Genus Isistius Gill, 1865 sides. The mesial cutting edge is convex and may be ser- Isistius triangulus (Probst, 1879) rated or smooth. The mesiolabial depression of the root is Material: 295 lower teeth. (Fig. 2-4) 244 N. Vialle et al.

This species is represented by numerous teeth, all belong- Fig. 2 1: Centrophorus aff. granulosus (scale: 1 mm), lower anteriorc ing to the lower jaw. All the dental files are represented, tooth, a: labial view; b: lingual view. (MAZ 1); 2–3: Deania aff. calceus (scale: 1 mm), lower anterior tooth, a: labial view; b: lingual view; 3: from the anteriors to the commissurals. The anterior and upper tooth, labial view. (MAZ 2; MAZ 3); 4: Isistius triangulus (scale: antero-lateral teeth are very compressed labiolingually and 1 mm), commissural teeth, a: labial view; b: lingual view. (MAZ 4); 5: have a very large triangular symmetric crown. The mesial ‘‘Oligodalatias’’ sp. (scale: 1 mm), lower anterior tooth, lingual view. cutting edge is thin, translucent and weakly serrated. On (MAZ 5); 6: Squaliolus schaubi (scale: 1 mm), lower anterior teeth, a: lingual view; b: labial view. (MAZ 6); 7: Pristiophorus suevicus (scale: the labial face, the apron is made of a thin layer of enamel, 1 mm), rostral tooth, profile. (MAZ 7); 8: Squatina subserrata (scale: with a poorly marked limit. The layer ends behind a 1 mm), lateral tooth, a: labial view; b: lingual view. (MAZ 8); 9: Alopias ‘‘button-hole’’ which divides the root into two lobes. On aff. superciliosus (scale: 5 mm), lateral tooth, a: labial view; b: lingual the root, square or rectangular in shape, there is a foramen view. (MAZ 9); 10: Carcharoides catticus (scale: 5 mm), upper lateral tooth, a: lingual view; b: labial view. (MAZ 10); 11: Cosmopolitodus on the top of the ‘‘button-hole’’. The commissural teeth hastalis (scale: 5 mm), lateral tooth, a: labial view; b: lingual view. (Fig. 2-4) are asymmetrical and have a poorly defined (MAZ 11); 12: Mitsukurina lineata (Scale: 5 mm), anterior tooth, a: distal heel. The ‘‘button-hole’’ is very low and the external labial view; b: profile; c: lingual view. (MAZ 12) groove is oblique. Remarks. The Neogene species I. triangulus differs from the Palaeogene species I. trituratus Winkler 1874 by are smooth and have a convex shape. We also observe the the lower position of the ‘‘button-hole’’ on the labial face of presence of a well-developed distal heel. The apron is the root and the serrated cutting edges (Ledoux 1972). The broad, flat and spreads on all the surface of the labial face species I. triangulus is very common in the Miocene (Fig. 2-6b). The apron is separated into two unequal lobes deposits of north Hemisphere, including the south of on each side of the median labial foramen. The labial France from He´rault (Cappetta 1970) to Vaucluse (Ledoux depression of the root is well marked. On the lingual face 1972). (Fig. 2-6a), the crown-root boundary stops at the level of Genus ‘‘Oligodalatias’’ Welton, 1979 the relatively large median lingual foramen. The lingual ‘‘Oligodalatias sp.’’ depression is well developed but its edges are not well Material: 2 lower teeth. (Fig. 2-5) marked. The basal part of root lacks on our material. Remarks. The absence of lingual marginal foramina on The material consists of two poorly preserved teeth, the lingual face of root allows to assign these teeth to reduced to the crown. Very close to the genus Squaliolus, S. schaubi. This genus is very rare in the fossil record (Casier these teeth differ, however by a mesial and a distal cutting 1958, 1966; Barthelt et al. 1991) and was only found in edge strongly serrated. On the distal side there is a heel Palaeogene and Neogene deep-water deposits. This species separated to the main cusp by a notch. was reported in the deposits of the Early and Middle Remarks. This genus was described for the first time by Miocene of southern France (Cappetta 1970; Ledoux Welton (1979) with two species: ‘‘Oligodalatias’’ jordani 1972). from the Middle Oligocene of the Pittsburg Bluff Formation (Colombia County, Oregon, USA) and ‘‘Oligodalatias’’ roe- Order Pristiophoriformes Berg, 1958 deri from the Middle Eocene of the Ardath Formation (San Family Pristiophoridae Bleeker, 1859 Diego County, California, USA). If the oldest species Genus Pristiophorus Mu¨ller & Henle, 1837 Oligodalatias roederi was recently synonymised to Squa- Pristiophorus suevicus (Jaekel, 1890) liodalatias weltoni (Adnet et al. 2006), our material is remi- Material: 1 rostral tooth. (Fig. 2-7) niscent of the second one: ‘‘Oligodalatias’’ jordani. Even if it This rostral tooth is long and thin with a sword shape. It is an unpublished nomen nudum of Welton (1979), we refer consists of a basal peduncle and a long, flat cap more or the teeth of Mazan to this genus, pending supplementary less curved to the rear. The peduncle is slightly smaller material. This invalid genus was already recorded from the than the crown. The cap is translucent and reveals Aquitanian of southern France by Cappetta (1970,plate7, the internal nutritive ducts. The base of the peduncle is Fig. 8-9) as an indeterminate Squaliformes and by Ledoux wider than the top, and in basal view there is a deep con- (1972, Fig. 12) as belonging to a new species of Squaliolus. cavity in the form of funnel which contains the pulp of the Genus Squaliolus Smith & Radcliffe, 1912 tooth. Squaliolus schaubi (Casier, 1958) Remarks. P. suevicus, recovered from the Miocene of Centroscymnus schaubi (Casier, 1958) southern Germany (Jaekel 1890; Barthelt et al. 1991), is Material: 3 lower teeth. (Fig. 2-6) uncommon in the Miocene sediments of southern France (Cappetta 1987). Nevertheless, it was also recorded from a In this genus, teeth have a crown with a cusp strongly bent rostral tooth, in the Middle Miocene deposits of ‘‘Cabrie`res towards the rear. The mesial and the distal cutting edges d’Aigues’’ (Brisswalter 2009). A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) 245 246 N. Vialle et al.

Order Squatiniformes Buen, 1926 The fossil remains of this genus are essentially represented Family Squatinidae Bonaparte, 1838 by isolated fragments of gill-rakers. These gill-rakers are Genus Squatina Dumeril, 1806 laterally flattened, curved with a hatchet-shaped basal part Squatina subserrata Mu¨nster, 1846 extending by a long and slender rod. These fragments are Material: 1 tooth. (Fig. 2-8) entirely covered by enameloid. Remarks Our remains may be attributed to the Oligo- The single tooth is poorly preserved. The crown, straight Miocene species C. parvus instead of the living species and high, has a triangular outline in labial view and is C. maximus Gunner 1765 as commonly accepted. perpendicular to the flattened root. The cusp is slightly bent Although this genus was first identified in the fossil lingually. The cusp presents at its base an enameled state based on gill-rakers, some oral teeth were found in expansion (Fig. 2-8a), forming a distinct rounded apron. ‘‘Cabrie`res d’Aigues’’ (Brisswalter 2009). The genus The lingual face (Fig. 2-8b) of the crown is more convex was also recorded from the Middle Miocene of ‘‘Bon- than the labial face. On each side of the cusp there are well- pas’’ (Cappetta et al. 1967). Cappetta and Ledoux developed heels. The cutting edges are sharp and extend on (1970) observed that the fragments of branchioctenii the two heels. The root is very spread lingually and pre- collected in the Mediterranean fossil deposits are usu- sents a well-marked protuberance. At the junction between ally those of young individual as are those collected at the crown and the protuberance of the root, there are ‘‘Mazan’’. numerous small foramina. The basal face of the root is flat with a well-marked median triangular depression. Family Lamnidae Mu¨ller & Henle, 1838 Remarks. If this species is well recorded in the Miocene Genus Carcharoides Ameghino, 1901 of Europe and North America, this taxon is relatively Carcharoides catticus (von Philippi, 1846) scarce in southern France. However, it was recorded by Otodus catticus von Philippi, 1846 Cappetta (1973, 1987) in the Early Burdigalian of ‘‘Lespi- Lamna cattica, in Cappetta, 1970 gnan’’ (He´rault) and in the Middle Miocene of ‘‘Cabrie`res Material: 4 teeth. (Fig. 2-10) d’Aigues’’ (Brisswalter 2009). These teeth are characterized by a high cusp, large at its Order Lamniformes Berg, 1958 base and bent toward the rear. The cutting edges extend on Family Alopiidae Bonaparte, 1838 the entire height of the main cusp and are smooth. On both Genus Alopias Rafinesque, 1810 sides of the main cusp we note the presence of a well Alopias aff. superciliosus (Lowe, 1840) developed, high, strong and sharp accessory cusplet. The Carcharias superciliosus Lowe, 1840 mesial cusplet is bigger and straighter than the distal cus- Material: 1 tooth. (Fig. 2-9) plet, which bents toward the rear like the main cusp. The two cusplets bear more or less marked cutting edges. The This very lateral tooth has a triangular, high and slender root is very thin and flat. On its basal face, there is the trace cusp, which overhangs the root in labial view. The crown of a groove. bends toward the rear. The cutting edge is more developed Remarks. It is the first evidence of this taxon in the on the distal side than on the mesial side. The labial face of Vaucluse area. It was previously reported from the Early crown is relatively flat contrary to the lingual face, which is Burdigalian of He´rault (Cappetta 1970) and elsewhere in much cambered. The base of the crown distinctly over- North Hemisphere during Oligocene (e.g. Mu¨ller 1983, hangs the labial face of the root, which is massive and 1999) and Miocene (e.g. Bollinger et al. 1995; Purdy et al. transversely elongated, with two well-defined lobes. The 2001). mesial lobe of the root is slightly longer than the distal lobe. The basal face bears a poorly marked groove. Genus Cosmopolitodus Glu¨ckman, 1964 Remarks. This off-shore living species is known since Cosmopolitodus hastalis (Agassiz, 1843) the Early Miocene (e.g. Case, 1980; Purdy et al. 2001) but Isurus hastalis Agassiz, 1843 appears in the Middle Miocene deposits of southern France Material: 1 tooth. (Fig. 2-11) (Cappetta 1987). Brisswalter (2009) described the dental This tooth is very stocky and has a low triangular crown. remains of a second extant species of Alopias, A. aff. The cusp has a relatively flat labial face and a clearly vulpinus Bonnaterre 1788 in ‘‘Cabrie`res d’Aigues’’ (Vau- convex lingual face. The cusp bends slightly toward the cluse, southern France). rear. The base of the cusp spreads marginally on the lobes Family Cetorhinidae Gill, 1862 of the root. The cutting edges of the cusp are completely Genus Cetorhinus Blainville and de, 1816 smooth and continue to the heels. The root is well devel- Cetorhinus parvus Leriche, 1908 oped and has a massive aspect with two expanded but Material: Some fragments of gill-rakers. poorly differentiated lobes. A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) 247

Remarks. Distributed in the Mio-Pliocene worldwide base of the crown to the half part of its height. This feature deposits, the occurrence of C. hastalis was previously depends of the specimen and the position on the jaw and recorded from He´rault (Cappetta 1970) and ‘‘Cabrie`res does not allow a specific separation (Antunes and Jonet d’Aigues’’ (Brisswalter 2009). 1970). Commissural teeth (Fig. 3-2) have a very stocky crown with or without lateral cusplet. The root is very Family Mitsukurinidae Jordan, 1898 large, sometimes larger than the crown, and has a basal flat Genus Mitsukurina Jordan, 1898 face with a deep groove. Mitsukurina lineata (Probst, 1879) Remarks. In 1912, Joleaud reported two species from Material: 288 teeth. (Fig. 2-12) Vaucluse, Odontaspis contortidens Agassiz 1843 and Chi- The anterior teeth have a slender and high crown, with a loscyllum fossile Probst 1879 which were partly assigned sigmoid shape in profile view (Fig. 2-12b). The labial face afterward to Odontaspis acutissima by Cappetta (1969, 1970). (Fig. 2-12a) is relatively flat or even slightly convex, while This worldwide species is very abundant in the Miocene the lingual face is strongly convex. The enameloid of the deposits of southern France (Brisswalter 2009). lingual face (Fig. 2-12c) is marked by strong folds on its Carcharias sternbergensis Reinecke, Moths, Grant & surface. These folds are parallel in the lower part of the Breitkreuz, 2005 crown and more flexuous toward the apex where they Odontaspis molassica Joleaud, 1912 disappear. The heels are very short and abrupt, with sec- Material: 11 teeth. ondary cusplets present or not. The root is well developed with two lobes. The lateral teeth have shorter cusps, with The teeth of this species differ from the previous one by a more developed oblique heels compared to the anterior thin and slender crown with a sigmoid profile. The crown is teeth. The root has lobes more spread laterally and a deep smooth on both sides. The cutting edges disappear in the groove on its basal face. There is often a pair of small lower third of the crown. The labial and the lingual face marginal cusplets. have a convex shape giving to the base a subcircular sec- Remarks. Originally described by Probst (1879) in the tion. The root is absent on our material. Early Miocene of SW Germany, this species is abundant in Remarks. Teeth assigned to Odontaspis molassica bathyal Miocene deposits in Vaucluse (Joleaud 1912; Probst 1879 in the southern Rhodanian basin (Joleaud Cappetta 1975, 1987) and other Alpine Foreland basins 1907, 1912) and in He´rault (Cappetta 1970) were recently (Cappetta 1987; Bollinger et al. 1995). synonymised with C. sternbergensis by Cappetta (2006). Family Odontaspididae Mu¨ller & Henle, 1839 Order Carcharhiniformes Compagno, 1973 Genus Carcharias Rafinesque, 1810 Family Carcharhinidae Jordan & Evermann, 1896 Carcharias aff. acutissima (Agassiz, 1843) Genus Carcharhinus Blainville, 1816 Odontaspis contortidens Agassiz, 1843 Carcharhinus priscus (Agassiz, 1843) Chiloscyllum fossili Probst, 1879 Sphyrna prisca Agassiz, 1843 Material: 33 teeth. (Fig. 3-1, 3-2) Carcharhias (Aprionodon) stellatus Probst, 1878 Carcharhias (Aprionodon) brevis Probst, 1878 The lateral teeth (Fig. 3-1) have a high and slender cusp, Carcharhinus (Hypoprion) lusitanicus Jonet, 1966 (in which is slightly bent toward the rear. The cusp is very part) broad at the base and overhangs the root on the labial face. Carcharhinus priscus, in Cappetta, 1970 The cusplets are well defined and more developed than Material: 60 teeth. (Fig. 3-3, 3-4) those of the anterior teeth. On the more lateral teeth, cusplets have a triangular shape and remind those of the genus The upper teeth (Fig. 3-4) have a narrow triangular crown. Carcharoides discussed previously. The root is very long The cusp is well developed and bent toward the rear. The with two well-developed lobes, separated from each other cutting edges are serrated only on their lower part. On each by a deep groove. The anterior teeth have a more side of the cusp there is a well-developed heel. The distal straightened and slender cusp flanked by small, sharp heel is sometimes separated from the cusp by a notch. Both cusplets. The cutting edge is well developed. The para- heels bear stronger serration than cusp. The root is flat and symphyseal teeth, much smaller than the laterals and the has a lingual protuberance bearing a deep clear groove. The anteriors, have a crown with a very pronounced sigmoid lower teeth (Fig. 3-3) have a narrower cusp, bent toward profile. On each side, there is a small cusplet. The very the inside of the mouth. The cutting edges are smooth, as massive root has fused lobes and a strong lingual protu- the two well-separated heels. The root is low and mesio- berance. On the lingual face of some lateral, anterior and distally elongated. parasymphyseal teeth, enameloid is marked by many Remarks. Teeth of Carcharhinus are very abundant in irregular folds, more or less salient, spreading from the the Miocene deposits of southern France with two species 248 N. Vialle et al. well represented, C. elongatus Leriche, 1910 (Leriche Fig. 3 1–2: Carcharias aff. acutissima (scale: 5 mm), 13: lateral c 1957; Cappetta 1970) and mainly C. priscus (Cappetta tooth, a: labial view; b: lingual view; 14: commissural tooth, lingual view. (MAZ 13; MAZ 14); 3–4: Carcharhinus priscus (scale: 5 mm), 3: 1969, 1970, 1987; Brisswalter 2009). Purdy et al. (2001) lower tooth, a: labial view; b: lingual view; 4: upper tooth, a: lingual view; claimed that teeth of C. priscus (pl. 13: Figs. 8-19) b: labial view. (MAZ 15; MAZ 16); 5: Galeocerdo aduncus (scale: described by Cappetta (1970) are in fact synonym of the 5 mm), anterior tooth, a: labial view; b: lingual view. (MAZ 17); 6: extant species C. brachyurus Gu¨nther, 1870. The specific Isogomphodon acuarius (scale: 1 mm), lateral tooth, a: labial view; b: lingual view. (MAZ 18); 7: Rhizoprionodon ficheuri (scale: 1 mm), status of these fossil forms is still sufficiently uncertain so lateral tooth, a: labial view; b: lingual view. (MAZ 19); 8–9: Pachys- our teeth are attributed to C. priscus pending further cyllium aff. dachiardii (scale: 1 mm), 8: anterior tooth, a: labial view; b: analysis. lingual view; 9: lateral tooth, a: occlusal view; b: lingual view. (MAZ 20; MAZ 21); 10-11: Scyliorhinus cf. joleaudi (scale: 1 mm), 10: lateral Genus Galeocerdo Mu¨ller & Henle, 1837 tooth, a: lingual view; b: labial view; 11: anterior tooth, a & b: labial view; Galeocerdo anduncus Agassiz, 1843 c: lingual view. (MAZ 22; MAZ 23); 12: Sphyrna arambourgi (scale: For the synonymy, see Cappetta (1970:50-51) 5 mm), lateral tooth, a: lingual view; b: labial view. (MAZ 24) Material: 7 teeth. (Fig. 3-5) The teeth possess a broad triangular and high cusp, which Genus Rhizoprionodon Whitley, 1929 leans toward the rear. The cutting edge is incised by small Rhizoprionodon ficheuri (Joleaud, 1912) serrations, and the teeth lack a mesial heel. The distal Carcharias (Physodon) ficheuri Joleaud, 1912 cutting edge is shorter and straight, marked by serrations. Physodon miocanenicus Jonet, 1966 The distal heel is separated from the crown by a deep notch Scoliodon dentatus Jonet, 1966 (in part) and presents a cutting edge with well-marked serration Scoliodon taxandriae Jonet, 1966 (in part) decreasing toward the rear. The labial face is flat whereas Material: 28 teeth. (Fig. 3-7) the lingual face is clearly convex. The root is thick and The teeth have a small size with a cusp bent toward the possesses a well-developed bulge. rear. The base of the crown is rather extended, particularly Remarks. This worldwide species was reported in the on the mesial edge, where the heel is not differentiated. Miocene deposits of the Rhone valley by Leriche (1906), The distal heel is high, unserrated with a rounded outline in Joleaud (1912) and Brisswalter (2009). In addition, Capp- labial view. On the labial face (Fig. 3-7a), the crown etta (1969, 1970, 1987) recorded this species from the slightly overhangs the root. The mesial cutting edge is contemporaneous He´rault deposits. convex and smooth. The root is low and presents on its lingual face, a protuberance where a distinct deep groove Genus Isogomphodon Gill, 1862 appears. Isogomphodon acuarius (Probst, 1879) Remarks. This species is relatively abundant in the Alopecias acuarius Probst, 1879 Middle and the Late Miocene of southern France (He´rault, Carcharhinus (Aprionodon) lerichei, in Jonet, 1966 Cappetta 1970; Vaucluse, Joleaud 1912; Brisswalter 2009). Aprionodon acuarius, in Cappetta, 1970 It also occurs, but more rarely in the Miocene of Belgium Material: 8 teeth. (Fig. 3-6) (Leriche 1926) and North America (Mu¨ller 1999). Teeth present a slight sigmoid curved crown. The Family Hemigaleidae Hasse, 1879 enameloid is totally smooth. The cusp leans slightly Genus Paragaleus Budker, 1935 toward the inside of the mouth. The cutting edges spread Paragaleus aff. pulchellus (Jonet, 1966) to the base of the crown and are completely smooth. The Galeorhinus pulchellus Jonet, 1966 heels are well developed, elongated and sub-horizontal Material: 1 tooth. and do not share any separation from cusp. The root is almost horizontal with two lobes very enlarged and The single example of this species is represented by a rounded extremities. On the median protuberance of the fragment of crown, probably from a lower tooth. The cusp lingual face of the root (Fig. 3-6b) a well-marked groove is erect and presents a concave mesial cutting edge. On its appears. distal side, there is a small sharp cusplet which composes Remarks. This species, common in Miocene of north the distal heel. The root is totally lacking. Atlantic and Mediterranean, was reported by previous Remark. Due to the poor quality of material, the iden- authors (Leriche 1906; Priem 1912) and by Cappetta tification of this species is uncertain. However, this species (1970) from the Miocene deposits of the Rhone Valley. was reported by Cappetta (1970) in the Middle Miocene of Cappetta (1970) described Isogomphodon caunellensis, He´rault and by Brisswalter (2009) in the Middle Miocene from the Late Burdigalian of the region of Montpellier of ‘‘Cabrie`res d’Aigues’’ (Vaucluse). Both figures show (He´rault) which differs from I. acuarius by larger teeth similar features which support the referral of our tooth to with a wider cusp. this species. A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) 249 250 N. Vialle et al.

Family Scyliorhinidae Gill, 1862 Fig. 4 1: Galeorhinus sp. (scale: 1 mm), antero-lateral tooth, labial c Genus Pachyscyllium Reinecke, Moths, Grant & Breitk- view. (MAZ 25); 2: Rhynchobatus pristinus (scale: 1 mm), lateral tooth, occlusal view. (MAZ 26); 3–4: Raja gentili (scale: 1 mm), 3: anterior reuz, 2005 male tooth (MAZ 27), labial view; 4: anterior female tooth, lingual view Pachyscyllium aff. dachiardii (Lawley, 1876) (MAZ 28); 5–6: Dasyatis probsti (scale: 1 mm), 5: female tooth (MAZ Scyliorhinus dachiardi Lawley, 1876 29), lingual view; 6: male tooth, lingual view. (MAZ 30): 7: Dasyatis Scyliorhinus distans Probst, 1879 (in part) rugosa (scale: 1 mm), female tooth, lingual view. (MAZ 31); 8–9: Taeniura aff. cavernosa (scale: 1 mm), 8: female tooth, lingual view. Premontreia dachiardi, in Cappetta, 2006 (MAZ 32); 9: male tooth, lingual view. (MAZ 33); 10: Mobula pectinata Material: 83 teeth. (Fig. 3-8, 3-9) (scale: 1 mm), a: occlusal view; b: basal view. (MAZ 34); 11: Aetobatus arcuatus (scale: 10 mm), upper tooth, occlusal view. (MAZ 35); 12: The main cusp is slender and straight with a sigmoid Myliobatidae indet. (scale: 10 mm), part of median tooth, a: occlusal profile. The lingual face is more convex than the labial. view; b: posterior view. (MAZ 36); 13: Plesiobatis sp. (scale: 10 mm), The cusp presents a transverse basal bulge overhanging the lingual view. (MAZ 37) root labially. This bulge bears sometimes parallel folds more or less marked (Fig. 3-9a). The cusp is slightly bent differentiated, but a clear distal heel is separated from the toward the inside of the mouth in lateral files. On each side cusp by a deep notch. The cutting edges are smooth. On the of cusp there is one pair of sharp and stocky cusplets. The labial face (Fig. 3-12b), the basal limit of the crown is flat. cutting edges are continuous from cusp to lateral cusplets. Remarks. In many previous works, there is confusion in The root presents a flat basal face and bears a strong lingual teeth assigned to Sphyrna that often belong to Carcharhi- protuberance. The root is divided by a deep groove. nus (Cappetta 1970, 1987). The genus Sphyrna is, however, Remarks. Reported by Cappetta (1970, 1987; Cappetta represented in the Middle Miocene of southern France by and Nolf 1991; Brisswalter 2009), this taxon is common in two species, the living Sphyrna zygaena Linnaeus, 1758 the Miocene deposits of southern France. (Cappetta 1970, 1987) and Sphyrna arambourgi (Cappetta Genus Scyliorhinus Blainville, 1816 1970; Brisswalter 2009) distinct from the previous one by Scyliorhinus cf. joleaudi Cappetta, 1970 smaller teeth and no serrated cutting edges. Material: 10 teeth. (Fig. 3-10, 3-11) Family Triakidae Gray, 1851 These teeth are small with a very large root. The main cusp Genus Galeorhinus Blainville, 1816 is very straight, slender and sharp, bending toward the rear Galeorhinus sp. and toward the inside of the mouth. The cutting edges are Material: 9 teeth. (Fig. 4-1) not well differentiated. The base of labial face (Fig. 3-10a) These small teeth are mainly represented by isolated of the crown largely overhangs the lobes of the root and crowns. The main cusp is high, bending toward the rear. bears small more or less marked folds. On each side of the The mesial cutting edge is slightly convex and not serrated. main cusp there are small accessory cusplets. The root is The distal heel is high and well developed and bears strong bulky with a massive lingual protuberance. The very spread cusplets decreasing in size toward the distal edge. The first basal face of the root does not present a groove. The lingual accessory cusplet is well separated from the main cusp. protuberance bears a small medio-lingual foramen. Remarks. Two species, G. affinis Probst, 1878 and Remarks. Some of our teeth (Fig. 3-11) are slightly G. latus Storms, 1894, were reported by Cappetta (1970)in different from the others by their labial smooth and flat face the Miocene of He´rault. Currently the first species is assigned as the material type of S. joleaudi. This morphology, dis- to Chaenogaleus affinis Probst, 1879 and the second one covered in the Langhian deposits of He´rault (Cappetta to Physogaleus latus Storms, 1894 (see Cappetta 2006). 1970) also occurs in the site of ‘‘Cabrie`res d’Aigues’’, More recently, Brisswalter (2009) reported the presence of Vaucluse (Brisswalter 2009) and in Portugal (Antunes et al. G. goncalvesi Antunes et al. (1999a) at ‘‘Cabrie`res d’Aigues’’. 1999b). These are currently considered as resulting from However, the very fragmentary remains of Mazan do not intraspecific variation in tooth morphology, as observable permit a reliable specific determination. in extant Scyliorhinus. Order Rajiformes Berg, 1940 Family Sphyrnidae Gill, 1872 Family Rhynchobatidae Garman, 1913 Genus Sphyrna Rafinesque, 1810 Genus Rhynchobatus Mu¨ller & Henle, 1837 Sphyrna arambourgi Cappetta, 1970 Rhynchobatus pristinus (Probst, 1877) Scoliodon taxandriae Jonet, 1966 (in part) Pristis pristinus Probst, 1877 Material: 5 teeth. (Fig. 3-12) Material: 31 teeth. (Fig. 4-2) The teeth are labio-lingually flattened with a triangular The teeth have a massive appearance. The oral face of the cusp bending toward the rear. The mesial heel is not crown may be differentiated into three different areas. The A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) 251 252 N. Vialle et al. labial area is more or less convex. The occlusal has a tri- root is composed by two narrow lobes separated by a deep angular shape and is separated from the labial one by a groove. In male teeth (Fig. 4-6), the posterior part of the clear straight crest. These two areas present a very precise crest stretches as a peak, the medio-external hollow is la- ornamentation of enameloid granules. The lingual face of biolingually stretched, the ornamentation is coarse and the the crown is smooth and bears a well-developed enameled crest bears strong folds. protuberance forming a long central uvula. The root is Remarks. D. probsti differs from Taeniura cavernosa by massive with two lobes well separated by a deep groove the shape of the median labial hollow which is sub-circular with two centrally positioned foramina. On the lingual face for the first and triangular for the second. D. probsti is of the root, and on each side of the uvula, there is a well- recorded in the Langhian of He´rault (Cappetta 1970) and marked depression with a big margino-lingual foramen. was also mentioned by Brisswalter (2009) from the Ser- Remarks. The teeth of R. pristinus are relatively abun- ravallian of ‘‘Cabrie`res d’Aigues’’ (Vaucluse, southern dant in the Miocene deposits of He´rault (Cappetta 1970) France). and Vaucluse (Joleaud 1912; Brisswalter 2009) and elsewhere (e.g. Case 1980; Bollinger et al. 1995;Mu¨ller 1999). Dasyatis rugosa (Probst, 1877) Raja rugosa Probst, 1877 Family Rajidae Bonaparte, 1831 Material: 7 teeth. (Fig. 4-7) Genus Raja Linnaeus, 1758 Raja gentili Joleaud, 1912 On female teeth (Fig. 4-7), the crown is low and has a Material: 16 teeth. (Fig. 4-3, 4-4) transversal crest, broad and smooth, which overhangs the latero-posterior face. The labial face presents a narrow and These teeth are very small and we note on this taxa a strong extended laterally median hollow and has a reticulated sexual heterodonty. The males (Fig. 4-3) possess teeth with ornamentation. The anterior visor is marked, on its upper a high and slender cusp. The labial face of the cusp is part, by a bulge with a narrow groove. The root possesses marked by a slightly sharp cutting edge. The crown over- two large lobes with a triangular shape in basal view. The hangs the labial face of the root with a round-contoured and teeth of males have a pointed crest bent toward the rear. well-developed visor. The root is high with two large lobes The labial face is flat to slightly convex. It bears a more or separated by a small groove. The females (fig. 4-4) possess less medial hollow and a reticulated ornamentation. The rounded teeth with a pyramidal crown. Both face of the posterior visor has a concavity at the level of the medio- crown are smooth. The crown is divised by a transversal lingual ridge. median crest. On the lingual face, the uvula is not well Remarks. This species was described by Probst (1877) marked. The root is massive with two well-defined lobes, from the Early Miocene of SW Germany. Since, it was separated by a deep groove. recorded by Cappetta (1970) and Brisswalter (2009) from Remarks. As the living Rajidae, preferentially inhabit the Miocene of He´rault and Vaucluse, respectively. the deep sea bottom in warm temperate and tropical realms, their fossils are not frequent in the coastal Miocene Genus Taeniura Mu¨ller & Henle, 1837 deposits of southern France. However, this species was Taeniura aff. cavernosa (Probst, 1877) recognized in some neritic deposits in He´rault (Cappetta Raja cavernosa Probst, 1877 1970) but especially in the bathyal faunas of Vaucluse Dasyatis carvernosa, in Cappetta, 1970 (Joleaud 1912; Cappetta et al. 1967; Brisswalter 2009). Material: 21 teeth. (Fig. 4-8, 4-9) Order Myliobatiformes Compagno, 1973 Female teeth (Fig. 4-8) have a slightly convex labial face Family Dasyatidae Jordan, 1888 in profile view. The median hollow is labialy bordered by a Genus Dasyatis Rafinesque, 1810 distinct crest. This depression is deeper labially than lingually. The root is high with flattened lobes separated by a Many teeth of this genus occur in the deposits of Mazan broad groove. The teeth of males are cuspidate (Fig. 4-9). and they are often badly preserved. Nevertheless, some rare Remarks. In the fossil record this genus are often con- teeth could be attributed to previously described species. fused Dasyatis. Teeth of Taeniura differ from Dasyatis by Dasyatis probsti Cappetta, 1970 a sharp transversal edge and an occlusal face quite con- Material: 6 teeth. (Fig. 4-5, 4-6) cave. First recognised from the Messinian of Portugal (Antunes et al. 1999b), this genus appears relatively com- The female teeth (Fig. 4-5) have a big size with a crown mon in the Mediterranean deposits from the Langhian higher than the root. The transversal crest is broad and (Cappetta and Cavallo 2006). high. The median labial hollow has a sub-circular shape surrounded by a well-marked sharp edge. The lingual visor Family Mobulidae Gill, 1893 overhangs clearly the notch and the lobes of the root. The Genus Mobula Rafinesque, 1810 A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) 253

Mobula pectinata Cappetta, 1970 & Hovestadt-Euler, 1999 Pteromylaeus Garman, 1913 or Material: 2 teeth. (Fig. 4-10) Aetomylaeus Garman, 1908. The occlusal face (Fig. 4-10a) of the crown slopes toward Family Plesiobatididae Nishida, 1990 the front. Its surface is bumpy. The anterior visor is well Genus Plesiobatis Nishida, 1990 developed and widely overhangs the anterior face of the Plesiobatis sp. root. The transversal crest is cut out by long and pointed Material: 7 teeth. (Fig. 4-13) indentations directed backward. The lingual face of the The teeth have a globular shape, with a labial face sepa- crown has a concave profile. The posterior visor is less rated from the lingual face by a smooth crest. The labial developed than the labial one. The root has a flat basal face face presents a slightly reticulated ornamentation. A simi- (Fig. 4-10b) and is composed by four lobes. lar ornamentation can be observed on the apical half of the Remarks. This fossil species of pelagic ray was descri- lingual face. The lower part of the labial visor is very flat bed for the first time in the Langhian of He´rault (Cappetta and the visor is very sharp in profile view. The root is well 1970) like the other species M. loupianensis Cappetta, developed but not very thick, with two lobes spread la- 1970. However, the species observed in Mazan differs from biolingually and separated by a broad groove with a fora- the last (M. loupianensis) by the presence of a multi-cusped men. The basal face of the lobes is broad and very flat. transversal crest on tooth crown. It is the first occurrence of Remarks. The lack of a well-marked transversal crest M. pectinata in the Miocene deposits of Vaucluse. and the presence on the lingual face of Plesiobatis of a Family Myliobatidae Bonaparte, 1838 well-marked mediolingual crest extending from the trans- Genus Aetobatus Blainville, 1816 versal crest to the posterior edge of the lingual visor allow Aetobatus arcuatus (Agassiz, 1843) differentiating Plesiobatis to Dasyatis, and especially to Aetobatis arcuatus Agassiz, 1843 D. rugosa recovered in the deposit. Teeth of this genus Material: 2 fragmentary teeth. (Fig. 4-11) have been found in the Langhian of southern France (Cappetta 2006). The fragments are those of upper teeth. The teeth are fairly rectilinear except the lateral edges which are curved towards the back. The length of the crown decreases Paleoenvironmental inferences toward the lateral extremities. The labial face of the teeth is sub-vertical while the lingual face is oblique. The separa- All the fossiliferous deposits of Vaucluse analysed tion between the crown and the root is made by a well- here (‘‘Saint Didier’’, ‘‘Bonpas’’, ‘‘Caumont’’, ‘‘Cabrie`res marked posterior bulge. The root bends lingually and is d’Aigues’’ and ‘‘Mazan’’) are coeval. Faunal differences composed of a succession of grooves and laminae. The root are interpreted here as purely ecological. The new data is higher than the crown in the median region of the file. provided by the ‘‘Mazan’’ fauna, together with those of the Remarks. The dental plate of this genus is composed by other Middle Miocene localities of Vaucluse previously a single dental file. This fossil species of eagle ray is rel- reported in literature (Table 1) provides the first overview atively common in the Early and Middle Miocene of of the diversity of selachian fauna inhabiting the southern He´rault (Cappetta 1970) and Vaucluse (Cappetta et al. Rhodanian basin of the Mesogean realm during the trans- 1967; Brisswalter 2009). gressive cycle of the Langhian (see Besson et al. 2005). Based on the fauna from similar Vaucluse deposits, Myliobatidae indet. Cappetta et al. (1967) proposed an estimate of the sea level. Material: Many fragments. (Fig. 4-12) Authors estimated that the sea have reached a depth of The crown has hexagonal shape in occlusal view. The between 200 and 300 m at the Middle Miocene. The median file teeth are broader than long and generally rec- presence of many fossil representatives of extant pelagic tilinear. The surface of the crown (Fig. 4-12a) is flat and is and bathyal genera of selachian at ‘‘Mazan’’ (see Table 1); slightly convex in profile. On the lingual face (Fig. 4-12b), including the bathyal Squaliformes confirms the previously there is a well-developed bulge separating the crown from estimated depths. According to conclusion of Cappetta the massive root, which is composed by a succession of et al. (1967), the presence of numerous taxa as Aetobatus laminae and grooves in basal view. The lateral teeth have a arcuatus and many Carcharhinidae at Mazan is character- hexagonal shape like the median teeth, but they are as long istic of a quite warm water fauna. as broad. In terms of diversity, the selachian fauna of ‘‘Mazan’’ is Remarks. The very fragmentary state of our material close to that of ‘‘Cabrie`res d’Aigues’’ reported by Bris- precludes referral to a definite species and genera as swalter (2009) in the south of the Luberon Park (southern Myliobatis Cuvier (ex Dumeril), 1816, Weissobatis Hovestdat Vaucluse). Of the 40 species reported at ‘‘Cabrie`res 254 N. Vialle et al.

Table 1 Fossil selachians recovered in Mazan and comparison with the other Middle Miocene localities of Vaucluse (4) Saint Didier (2) Caumont (1) Bonpas (3) Mazan (5) Cabrie`res d’Aigues Habitat of living representatives

Paraheptranchias repens X X 27–1,000* Hexanchus sp. X X 0–1,875 Centrophorus aff. granulosus X X X X 50–1,440 Deania aff. calceus X X X X 70–1,470 Isistius triangulus X X X X 0–3,500 ‘‘Oligodalatias’’ sp. X X 0–4,000* Squaliolus schaubi X X X 200–2,000 Pristiophorus suevicus X X X 100–952 Squatina subserrata X X X 0–494 Alopias aff. superciliosus X X X 0–732 Cetorhinus parvus X X 0–904 Carcharoides catticus X 0–1,000* Cosmopolitodus hastalis X X 0–1,300* Mitsukurina lineata X X 270–960 Carcharias aff. acutissima X X X 0–191 Carcharias sternbergensis X X 0–191 Carcharhinus priscus X X 0–430 Galeocerdo anduncus X X X 0–140 Isogomphodon acuarius X X 4–40 Rhizoprionodon ficheuri X X 1–100 Paragaleus aff. pulchellus X X 0–100 Pachyscyllium aff. dachiardii XX 0–[2,000* Scyliorhinus cf. joleaudi X X 0–754 Sphyrna arambourgi X 0–275 Galeorhinus sp. X 2–471 Rhynchobatus pristinus X X X 0–65 Raja gentili X X X 3–1,000 Dasyatis probsti X X 0-480 Dasyatis rugosa X X 0–480 Taeniura aff. cavernosa X 0–500 Mobula pectinata X 0–100 Aetobatus arcuatus X X X 0–100 Myliobatidae indet. X X X 0–200 Plesiobatis sp. X X 44–680 For Caumont, Bonpas and Saint-Didier, Ledoux, 1972; for Bonpas, Cappetta et al. 1967; for Cabrie`res d’Aigues, Brisswalter, 2009 The maximal depth range (in meter) of living representatives (species or genus) is compiled from Compagno et al. 2005; Kyne and Simp- fendorfer 2007 and online data available at www.fishbase.org. * indicates that maximal depth range is that of the family d’Aigues’’, 23 are present in Mazan (66% taxa in com- Lamniformes (e.g. Alopias superciliosus, Cetorhinus par- mon). Most of them belong to the order of Carcharhini- vus, Cosmopolitodus hastalis) or pelagic ray (e.g. Mobula) formes (e.g. Carcharhinus priscus, Galeocerdo aduncus, testify of a more open environment than at ‘‘Cabrie`res Isogomphodon acuarius or Scyliorhinus joleaudi). Living d’Aigues’’. The site of ‘‘Bonpas’’ near Avignon, published representatives of those forms live in neritic to pelagic by Ledoux (1972), has 16 species in common (48%) environments (see Table 1) and indicates a shallow area with ‘‘Mazan’’. The main pool of taxa in common with close to the coastal zone (Brisswalter, 2009). Six taxa ‘‘Mazan’’ belongs to the order of Squaliformes with Cen- of batoid are newly reported compared to ‘‘Cabrie`res trophorus granulosus, Deania calceus, ‘‘Oligodalatias’’, d’Aigues’’ (e.g. Rhynchobatus pristinus, Dasyatis rugosa Isistius triangulus and Squaliolus schaubi. The presence of or Aetobatus arcuatus) and their living representatives are these forms is clearly characteristic of a relatively deep-sea distinctive of shallow water. The presence of pelagic environment (see Table 1) as indicated by the ecology of A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) 255 living representatives of C. granulosus or D. calceus that deposits of Vaucluse (see Table 1), we can propose the spend the majority of their life cycle at depths below 200 m environmental settings of deposit formation in ‘‘Mazan’’. (Kyne and Simpfendorfer 2007). Some of these taxa are These deposits correspond to those of an external platform also found in the deposits of ‘‘Caumont’’ and ‘‘Saint and/or upper slope, relatively open with a great depth Didier’’ (Vaucluse) considered as deposited in deep-sea influence. Presence of teeth of the most neritic inhabitants environment (Ledoux 1972). For instance, the Lamnifor- could have been transported far offshore within the sub- mes Mitsukurina lineata, known at ‘‘Saint-Didier’’, is a marine valleys. good indicator of deep-sea deposits because its living Looking at the faunal association of the site of ‘‘Cab- representative M. owstoni only inhabits the bathyal sea rie`res d’Aigues’’, which also contains a significant assem- bottom deeper than 200 m (Compagno 2001). Other newly blage, we note a similar grouping of dominant orders with recorded taxa from ‘‘Mazan’’ are in agreement with a deep- the same proportions; Carchariniformes, 31%; Lamnifor- sea origin of deposit (see Table 1)asParaheptranchias mes, 17%; Myliobatiformes, 19% and Squaliformes, 13%. repens, Hexanchus sp., Squaliolus shaubi, Pristiophorus This ecological similarity suggests that the environmental suevicus, Raja gentili and Plesiobastis sp. (known at settings in ‘‘Mazan’’ were probably more similar with ‘‘Bonpas’’ too, Cappetta, pers. obs.). Finally, three groups ‘‘Cabrie`res d’Aigues’’ than with the nearest localities dominate in ‘‘Mazan’’ deposits. The most important is the (‘‘Bonpas’’, ‘‘Caumont’’ and ‘‘Saint-Didier’’) considered as Carcharhiniformes with nine identified species, represent- deposited in strict deepwater environments. Such spatial ing 27% of species present at the site. The other two groups heterogeneities in the Middle Miocene selachian associa- are Lamniformes and Myliobatiformes, with seven species tions supply further arguments to consider that the conti- each, representing 20% of the association. These groups, nental shelf forming the southern Rhodanian basin during comprising lot of epipelagic or epibenthic representatives the Langhian was probably crenulated by numerous sub- today, support the idea of a relatively open sea with a near marine canyons or valleys. shore. On the other hand, the presence of numerous The new data provided by the fauna of ‘‘Mazan’’ allow a Squaliformes (i.e. 5 species constituting 14% of species in better understanding on the environment of the southern the site), Hexanchiformes, Rajidae and fossil representativs Rhodanian basin during the Middle Miocene but it would of Mitsukurina suggest a deeper environment, as in ‘‘Saint- be interesting to better explore Miocene deposits of the Didier’’, ‘‘Bonpas’’ or ‘‘Caumont’’. Vaucluse and adjacent areas in order to more precisely Such ecological incongruence could be explained by the redraw the outlines of Middle Miocene coasts. peculiar topology of the ancient seafloor of this area. Besson et al. (2005) clearly demonstrated that the Langhian Acknowledgments Authors are particularly grateful to the two sandy marls (‘‘Schlier’’ marls Formation) fill a large anonymous reviewers who significantly improved the quality of the first draft, and to Isabelle Vea for her corrections of English. complex network of incised valleys resulting from deep fluvial incisions of the Rhodanian basin during the regressive cycle of the Late Burdigalian (see Fig. 1). The References fossiliferous site of ‘‘Mazan’’ is precisely located near one of these submarine valleys. In such topological context, Adnet, S., Cappetta, H., & Reynders, J. (2006). Nouveaux genres de two alternatives hypothesis can explain why the fossil Squaliformes (Chondrichthyes) du Paleóge`ne des Landes (Sud- remains belonging to two different ecological associations Ouest de la France). Palaöntologische Zeitschrift, 80(1), 60–67. could be mixed in the same place: either the deep water Agassiz, L. (1843). Recherches sur les poissons fossiles. 3, 390 ? 32 P., 47 pl. faunal elements were migrating up submarine canyons into Ameghino, F. (1901). L’aˆge des formations se´dimentaires de shallow water by coastal upwelling, or the shallow water Patagonie. Anales de la Sociedad Cientı´fica Argentina, 51, elements were being washed into deeper water by turbidity 20–39 ? 65–91. currents or slumping. No fine data concerning the sedi- Antunes, M. T., Balbino, A. C., & Cappetta, H. (1999a). A new shark, Galeorhinus goncalvesi nov. sp. (Triakidae, Carcharhiniformes) mentology or tectonic are currently available to resolve from the latest Miocene of Portugal. Tertiary Research, 19(3–4), definitively this question. However, the dental elements of 101–106. the more neritic taxa are more badly conserved compared Antunes, M.T., Balbino, A.C. & Cappetta, H. (1999b). Se´laciens du to those of the deeper representatives. This suggests that a Mioce`ne terminal du bassin d’Alvalade (Portugal) Essai de synthe`se. Cieˆnsas da Terra 13, 115–129. minor part of material (e.g. Rhizoprionodon, Paragaleus, Antunes, M. T., & Jonet, S. (1970). Requins de l’Helve´tien supe´rieur Rhynchobatus, some Dasyatidae and Myliobatidae) may et du Tortonien de Lisbonne. Bulletin de l’Universite´ de have been slightly reworked and transported into deeper Lisbonne, 2, 120–280. waters. Barnard, K. H. (1925). A monography of the marine fishes of South Africa. Part I. Annals of the South African Museum, 21, 1–418. Starting from the faunal combination of the site of Barthelt, D., Fejfar, O., Pfeil, F. H., & Unger, E. (1991). Notizen zu ‘‘Mazan’’ and data supplied from previous works on the einem Profil der Selachier Fundstelle Walbertsweiler im Bereich 256 N. Vialle et al.

des mioza¨nen Oberen Meeresmolasse Su¨ddeutschlands. Mu¨nch- Cappetta, H., & Cavallo, O. (2006). Les Se´laciens du Plioce`ne de la ner Geowissenschaftliche Abhandlungen, 19, 195–208. re´gion d’Alba (Pie´mont, Italie Nord-Ouest). Rivista Piemontese Berg, L. S. (1940). Classification of fishes both recent and fossil. di Storia Natura, 27, 33–76. Trudy Zoological institut, Leningrad, 5, 85–517. Cappetta, H., Granier, J., & Ledoux, J.-C. (1967). Deux faunes de Berg, L.S. (1958). System der rezenten und fossilen Fischartigen und Se´laciens du Mioce`ne me´diterraneén de France et leur signifi- Fische (p. 310) Berlin: Deutsch Verlag Wiss. cation bathyme´trique. Comptes rendus sommaires des seánces de Besson, D. (2005). Architecture du bassin Rhodano-provençal la Socie´te´ geólogique de France, 7, 292. mioce`ne (Alpes, SE France). Relations entre de´formation, Cappetta, H. & Ledoux, J.C. (1970). Comparaison de la faune physiographie et se´dimentation dans un bassin molassique ichthyologique mioce`ne avec la faune actuelle de Me´diterraneé. d’avant-pays. (p. 449) Unpublish MSc thesis, The`se de l’Ecole Journeé ichthyologique C.I.E.S.M., Rome, pp. 21–23. des Mines-Paris Tech, Paris. Cappetta, H., & Nolf, D. (1991). Les se´laciens du Plioce`ne infe´rieur Besson, D., Parize, O., Rubino, J. L., Aguilar, J. P., Aubry, M. P., de Le-Puget-sur-Argens (Sud-Est de la France). Palaeonto- Beaudoin, B., et al. (2005). Un re´seau fluviatile d’aˆge Burdig- graphica A, 218, 49–67. alien terminal dans le Sud-Est de la France: remplissage, Case, G. R. (1980). A selachian fauna from the Trent Formation; extension, aˆge, implications. Compte rendu de Geoscience, lower Miocene (Aquitanian) of eastern North Carolina. Pala- 337, 1045–1054. eontographica, A, 171, 75–103. Blainville, H.M.D. de (1816). Prodrome d’une nouvelle distribution Casier, E. (1958). Contribution a` l’e´tude des poissons fossiles des syste´matique du re`gne animal. (pp. 105–112 ? 121–124) Bul- Antilles. Schweizerische palaeontologische Abhandlungen, 74, letin de la Socie´te´ Philomathique, Paris 8. 1–95. Bleeker, P. (1859). Over eenige vischsoorten van de Zuidkust- Casier, E. (1966). Faune ichthyologique du London Clay. wateren van Java. Natuurkundig Tijdschrift voor Nederlandsch XIV ? 403p. London: Trust. British Museum (Natural History). Indie¨,19, 329–352. Compagno, L.J.V. (1973). Interrelationship of living elasmobranchs. Bloch, M.E. & Scheinder, J.G. (1801). M.E. Blochii Systema In P.H. Greenwood, R.S., R.S. Miles & C. Patterson (Eds.) Ichthyologiae iconibus ex illustratum. Post obitum auctoris opus Interrelationships of Fishes. Zoological Journal of the Linnaean inchoatum absolvit correxit, interpolativ J.G. Scheinder, Saxo. Society, Supplement no.1–53, 15–98. pp. 584. Compagno, L.J.V. (2001). Sharks of the world. An annotated and Bollinger, T., Kindlimann, R., & Wegmu¨ller, U. (1995). Die marinen illustrated catalogue of shark species known to date. Volume 2. Sedimente (ju¨ngere OMM, St.Galler-Formation) am Su¨dwe- Bullhead, mackerel and carpet sharks (Heterodontiformes, strand des Ho¨rnlischu¨ttung (Ostschweiz) und die palo¨kologische Lamniformes and Orectolobiformes). FAO Species Catalogue Interpretation ihres Fossilinhaltes. Eclogae Geologica Helvetica, for Fishery Purpose, No. 1 (Vol 2, pp. 1–8 ? 1–269), Rome. 88, 885–909. Compagno, L.J.V., M. Dando, & S. Fowler. (2005). A field guide to Bonaparte, C. L. (1831). Saggio di una distribuzione metodica degli the sharks of the world (pp. 1–368) London: Harper-Collins. animali vertebrati. Giornale Arcadico di Scienze Lettere ed Arti, Cuvier, G. (1816). Le re`gne animal distribue´ d’apre`s son organisation, 52, 155–189. pour servir de base a` l’histoire naturelle des animaux et d’intro- Bonaparte, C. L. (1838). Selachorum tabula analytica. Nuovi Annali duction a` l’anatomie compareé. Paris Poissons, 2, 104–351. Scienze naturali, Bologna, 2, 195–214. de Buen, F. (1926). Cata´logo ictiolo´gico del Mediterrańeo espanõl y Bonnaterre, J.P. (1788). Ichthyologie. Tableau encyclope´dique et de Marruecos recopilando lo publicado sobre peces de las me´thodique des trois re`gnes de la (pp. 215). Paris: Nature, costas mediterrańea y pro´ximas del Atlańtico (Mar de Espanã) pl. A–B ? 1–100. (p. 221). Madrid: Resultados de la Campanã internacional del BRGM (1975). Carte Geólogique au 1/50,000 n° 941, Carpentras. Instituto Espanõl de Oceanographıá. BRGM Editions, 2-7159-1941-7. Demarcq, G. (1970). Etude stratigraphique du Mioce`ne rhodanien. Brisswalter, G. (2009). Inventaire des Elasmobranches (requins, raies, Me´moires du B.R.G.M., 61, 115–159. chime`res) des de´poˆts molassiques du Sud-Luberon (Mioce`ne Dumeril, A.M.C. (1806). Zoologie Analytique ou Me´thode Naturelle supe´rieur), a` Cabrie`res d’Aigues (Vaucluse) France (pp. 1–100). de classification des animaux, rendue plus facile a` l’aide de Courriers scientifiques du Parc Re´gional du Lube´ron Hors Se´rie: tableaux synoptiques, Paris. Apt. Fischer, H. (1878). Note paleóntologique sur la molasse de Cucuron. Budker, P. (1935). Description d’un genre nouveau de la famille des Bulletin de la Socie´te´ Geólogique de France, 3e`me se´rie T-7, Carcharhinide´s. Bulletin du Museúm d’Histoire Naturelle. Paris, 221–222. 2(7), 107–112. Garman, S. (1908). New Plagiostomia and Chimopnea. Bulletin of the Cappetta, H. (1969). L’Ichthyofaune (Euselachii, Teleostei) mioce`ne Museum of Comparative Zoology, Harvard, 51, 249–256. de la re´gion de Montpellier (He´rault). pp. 291. Unpublish MSc Garman, S. (1913). The Plagiostomia (Sharks, Skates and Rays). thesis, The`se de Spećialite´ (Montpellier). Memoirs of the Museum of comparative zoology, Harvard, 36, Cappetta, H. (1970). Les se´laciens du Mioce`ne de la re´gion de 1–528. Montpellier. Palaeovertebrata Me´moire extraordinaire (pp. Gignoux, M. (1960). Geólogie stratigraphique (5th ed.) Paris: Masson 1–140). et Cie. Cappetta, H. (1973). Les se´laciens du Burdigalien de Lespignan Gill, T. N. (1862). Analytical synopsis of the order of Squali; and (He´rault). Geóbios, 6, 211–223. revision of the nomenclature of the genera. Annals of the Lyceum Cappetta, H. (1975). Les Se´laciens mioce`nes du midi de la France, of Natural History of New-York, 7, 371–408. re´partition stratigraphique et bathyme´trique (p. 90). 3e`me reúnion Gill, T. N. (1865). Synopsis of the eastern American sharks. annuelle des Sciences de la Terre, Montpellier. Proceedings of the Academy of Natural Sciences of Philadel- Cappetta, H. (1987). Mesozoic and Cenozoic Elasmobranchii, phia, 16(5), 258–265. Chondrichthyes II. 3B. In H.P. Schultze (Ed.), Handbook of Gill, T. N. (1872). Arrangement of the families of fishes, or classes Paleoichthyology (p. 193). Pisces, Marsipobranchii, and Leptocardii. Smithsonian miscella- Cappetta, H. (2006). Elasmobranchii post-Triadici (index generum et neous collections, 5(2), 1–49. specierum). In W. Riegraf (Ed.), Fossilium Catalogus I: Gill, T. N. (1893). Families and subfamilies of fishes. Memoirs of the Animalia 142 (p. 472). Backhuys Publish: Leiden. National Academy of Science, 6, 127–138. A new shark and ray fauna from the Middle Miocene of Mazan, Vaucluse (southern France) 257

Glu¨ckman, L.S. (1964). Class Chondrichthyes, Subclass Elasmobran- Leriche, M. (1908). Sur un appareil fanonculaire de Cetorhinus trouve´ chii. In D.V. Obruchev (Ed.), Fundamentals of Paleontology, a` l’e´tat fossile dans le Plioce`ne d’Anvers. Compte Rendu Academia Nauk SSSR (Russian edition) (Vol. 11, pp.196–237) Acade´mie des Sciences, Paris, 146, 875–878. (pp. 292–352 in English translation, Jerusalem, 1967). Leriche, M. (1910). Les poissons oligoce`nes de la Belgique. Goodrich, E.S. (1909). Vertebrata Craniata. I. Cyclostomes and fishes. Me´moires du Museé Royal d’Histoire Naturelle de Belgique, 5, In. E.R. LANKESTER (Ed.) A Treatise on Zoology (pp. 233–363. XVI ? 518), pt. 9, London. Leriche, M. (1926). Les poissons tertiaires de la Belgique. IV. Les Gray, J.E. (1851). List of the specimens of fish in the collection of poissons neóge`nes. Me´moires du Museé Royal d’Histoire the British Museum, Part 1. British Museum of Natural History Naturelle de Belgique, 32, 367–472. (pp. 160) Leriche, M. (1957). Les Poissons neóge`nes de la Bretagne, de la Gunner, J. E. (1765). Brugden (Squalus maximus), Beskrvenen ved. Anjou et de la Touraine. Me´moire de la Socie´te´ Geólogique de J.E. Gunnerus. Det Trondhei Sel’skoe Skrif, 3, 33–49. France N.S., 36, 1–61. Gu¨nther, A. (1870). Catalogue of the fishes in the British Museum, 8, Linnaeus, C. (1758). Systema Naturae, 10th edn (Vol. 1, pp. 230–338) Catalogue of the Physostomi containing the families Gymnoti- Nantes & Pisces. dae, Symbranchidae, Muraenidae, Pegasidae and of the Lopho- Lowe, R. T. (1839). A supplement to a synopsis of the fishes from branchii, Plectognathi, Dipnoi, Ganoidei, Chondropterygii, Madeira. Proceedings of the Zoological Society. London, 7(89), Cyclostomata (p. 549). London: Leptocardii in the collection 76–92. of the British Museum. Lowe, R. T. (1840). Description of certain new species of Madeiran Hasse, J. C. F. (1879). Das natu¨rliche System des Elasmobranchier fishes, with additional information relating to those already auf Grundlage des Baues und der Entwicklung ihrer Wirbelsaüle described. Proceedings of the Zoological Society, London, 8, (385 ? 27 pp). Jena: Gustav Fischer. 36–39. Hovestadt, D. C., & Hovestadt-Euler, M. (1999). Weissobatis Mu¨ller, A. (1983). Fauna und Palo¨kologie des marinen Mit- micklichi n. gen., n. sp., an eagle ray (Myliobatiforms, Mylio- teloligoza¨ns des Leipziger Tieflandsbucht (Bo¨hlener Schichten). batidae) from the Oligocene of Frauenweiler (Baden-Wu¨rttem- Altenburger Naturwissenschaftliche Forschungen, 2, 1–152. berg, Germany). Palaöntologische Zeitschrift, 73, 337–349. Mu¨ller, A. (1999). Ichthyofaunen aus dem atlantischen Tertia¨r des Jaekel, O. (1890). Ueber die Systematische Stellung und u¨ber fossile USA. Leipziger Geowissenschafteb, 9–10, 1–360. reste der Gattung. Pristiophorus Zeitschrift der Deutschen Mu¨ller, J., & Henle, F. G. J. (1837). Ueber die Gattungen der Geologischen Gesellschaft, 42, 86–120. Haifische und Rochen, nach ihrer Arbeit: ‘‘Uber die Naturge- Joleaud, L. (1907). Geólogie et Paleóntologie de la Plaine du Comtat schichte der Knorpelfische’’. Bericht Akademie der Wissens- et de ses abords. Description des terrains Neóge`nes. Me´moire de chaften zu Berlin, 1, 111–118. l’Acade´mie du Vaucluse, 1, 1–254. Mu¨ller, J., & Henle, F. G. J. (1838). On the generic characters of Joleaud, L. (1912). Geólogie et Paleóntologie de la Plaine du Comtat cartilaginous fishes, with descriptions of new genera. Magazine et de ses abords. Description des terrains Neóge`nes. Me´moire de of Natural History, new series, 2, 1–36. l’Acade´mie du Vaucluse, 2, 255–285. Mu¨ller, J. & Henle, F.G.J. (1839). Systematische Beschreibung der Jonet, S. (1966). Notes d’ichthyologie mioce`ne. II. Les Carcharhini- Plagiostomen, i-xxii ? 220 pp. Berlin. dae. Boletim do Museu e Laborato´rio Mineralo´gico e Geolo´gico Mu¨nster, G. G. (1846). Ueber die in der Tertia¨r-Formation des Wiener da Faculdade de Cieˆncias. Universidade de Lisboa, 10(2), Beckens vorkommender Fisch Ueberreste, mit Beschreibung 65–88. einiger neuen merkwu¨rdigen Arten. Beitra¨ge Petrefakten-Kunde, Jordan, D. S. (1888). Description of two new species of fishes from 7, 1–31. South America. Proceedings of the Academy of Natural Sciences Nishida, K. (1990). Phylogeny of the suborder Myliobatoidei. Mem. of Philadelphia, 39, 387–388. Fac. Fish. Hokkaido University, 37(1–2), 1–108. Jordan, D. S. (1898). Description of a species of fish (Mitsukurina Pfeil, F. H. (1981). Eine nektonische Fischfauna aus dem un- owstoni) from Japan, the type of a distinct family of Lamnoid teroligoza¨nen schno¨neker Fischschiefer des Galon Grabens in sharks. Proceedings of the California Academy of Sciences, 3(1), Oberbayern. Geologica Bavarira, 82, 357–388. 199–204. von Philippi, R.A. (1846). Tornatella abbreviata, Otodus mitis, Jordan, D. S., & Evermann, B. W. (1896). Check-list of the fishes and Otodus catticus und Myliobatis testae. Palaeontographica, 1(1), fish-like Vertebrates of North and Middle America. Rept. U.S. 23–26. Commnr Fish, 21, 207–584. Priem, F. (1912). Sur les poissons fossiles des terrains tertiaires Jordan, D. S., & Snyder, J. O. (1902). Descriptions of two news supe´rieurs du Sud de la France. Bulletin de la Socie´te´ Geólog- species of squaloid sharks from Japan. Proceedings of the United ique de France se´rie 4 t. XII, 213–245. States National Museum, 25, 79–87. Probst, J. (1877). Beitra¨ge zur Kenntniss der fossils Fische aus der Kyne, P. M. & Simpfendorfer, C.A. (2007). A collation and Molasse von Baltringer. II. Batoidei A. Gu¨nther. Klein-und summarization of available data on deepwater chondrichthyans: grosszahnige Rochen. Jahreshefte des Vereins fu¨r vaterla¨ndische biodiversity, life history and fisheries. IUCN SSC Shark Naturkunde in Wu¨rttemberg, 33, 69–103. Specialist Group for the Marine Conservation Biology Institute Probst, J. (1878). Beitra¨ge zur Kenntniss der fossils Fische aus der (pp. 1–137). Molasse von Baltringer. Hayfische (Selachoidei A. Gu¨nther). Lawley, R. (1876). Nuovi Studi sopra ai Pesci ed Vertebrati fossili Jahreshefte des Vereins fu¨r vaterla¨ndische Naturkunde in delle Colline toscane (p. 122). Wu¨rttemberg, 34, 113–154. Ledoux, J. C. (1972). Les Squalidae (Euselachii) mioce`nes des Probst, J. (1879). Beitra¨ge zur Kenntniss der fossils Fische aus der environs d’Avignon (Vaucluse). Documents des laboratoires de Molasse von Baltringer.Hayfische (Selachoidei A. Gu¨nther) geólogie de la faculte´ des sciences de Lyon, Notes et Me´moires, (Schluss). Jahreshefte des Vereins fu¨r vaterla¨ndische Naturkun- 52, 133–175. de in Wu¨rttemberg, 35, 127–191. Leriche, M. (1906). Re´vision de la faune ichthyologique des terrains Purdy, R.W., Schneider, V.P., Applegate, S.P., Mclellan, J.H., Meyer, neóge`nes du basin du Rhoˆne. In Association Française pour R.L. & Slaughter, B.H. (2001). The Neogene Sharks, Rays, and Avancement des Sciences. Compte rendu de la 35e`me session, Bony Fishes from Lee Creek Mine, Aurora, North Carolina. Lyon. Notes et Me´moires (pp. 335–352). Smithsonian Contribution to Paleobiology, 90, 71–202. 258 N. Vialle et al.

Rafinesque, C.S. (1810). Caratteru di alcuni nuovi generi e nueve Storms, R. (1894). Troisie`me note sur les poissons du terrain specie di animali (principalmente di pesci) e pinate della Sicilia, Rupe´lien. Bulletin de la Socie´te´ belge de Geólogie, de Paleón- con varie osservazioni sopra i medisimi (pp. 105), Palermo, tologie et d’Hydrologie, 8, 67–82. Italia. Welton, B.J. (1979). Late Cretaceous and Cenozoic Squalomorphii of Reinecke, T., Moths, H., Grant, A., & Breitkreuz, H. (2005). Die the NorthWest Pacific Ocean. pp. 53. Unpublished MSc thesis, Elasmobranchier des norddeutschen Chattiums, insbesondere des University of California Berkeley, Berkeley. Sternberger Gesteins (Eochattium, Obere Oligoza¨n). Palaeontos, Whitley, G. P. (1929). Additions to the Check-list of the Fishes of 8, 1–135. New South Wales. Australian Zoologist, 5(4), 353–357. Smith, H. M., & Radcliffe, L. (1912). The squaloid sharks of the Winkler, T.C. (1874). Me´moires sur des dents de poissons du terrain Philippine Archipelago. Proceedings of the United States bruxellien. Archives du Museé Teyler, 3(4), 295–256. National Museum, 41, 677–685.